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The Role of Microorganisms in the Formation of a Stalactite in Botovskaya Cave, Siberia – Paleoenvironmental Implications

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The Role of Microorganisms in the Formation of a Stalactite in Botovskaya Cave, Siberia – Paleoenvironmental Implications Biogeosciences, 10, 6115–6130, 2013 Open Access www.biogeosciences.net/10/6115/2013/ doi:10.5194/bg-10-6115-2013 Biogeosciences © Author(s) 2013. CC Attribution 3.0 License. The role of microorganisms in the formation of a stalactite in Botovskaya Cave, Siberia – paleoenvironmental implications M. Pacton1,*, S. F. M. Breitenbach1, F. A. Lechleitner1, A. Vaks2, C. Rollion-Bard3, O. S. Gutareva4, A. V. Osintcev5, and C. Vasconcelos1 1Geological Institute, ETH Zurich, Sonneggstrasse 5, 8092 Zurich, Switzerland 2Department of Earth Sciences, University of Oxford, South Parks Road, Oxford OX1 3AN, UK 3Centre de Recherches Pétrographiques et Géochimiques, UMR 7358, Université de Lorraine, 54500 Vandoeuvre-lès-Nancy, France 4Institute of the Earth’s Crust, Russian Academy of Sciences, Siberian Branch, 128 Lermontova Street, Irkutsk 664033, Russia 5Arabica Speleological Club, Mamin-Sibiryak Street, P.O. Box 350, Irkutsk 554082, Russia *now at: Laboratoire de Géologie de Lyon : Terre, Planètes, Environnement Campus de la Doua, Universite Lyon 1 2 Rue Raphael Dubois, 69622 Villeurbanne Cedex, France Correspondence to: M. Pacton ([email protected]) Received: 20 March 2013 – Published in Biogeosciences Discuss.: 8 April 2013 Revised: 25 July 2013 – Accepted: 15 August 2013 – Published: 27 September 2013 Abstract. Calcitic speleothems in caves can form through negative δ13C excursions. The microscale biogeochemical abiogenic or biogenic processes, or through a combina- processes imply that microbial activity has only negligible tion of both. Many issues conspire to make the assess- effects on the bulk δ13C signature in speleothems, which is ment of biogenicity difficult, especially when focusing on more strongly affected by CO2 degassing and the host rock old speleothem deposits. This study reports on a multiproxy signature. analysis of a Siberian stalactite, combining high-resolution microscopy, isotope geochemistry and microbially enhanced mineral precipitation laboratory experiments. The contact between growth layers in a stalactite ex- 1 Introduction hibits a biogenic isotopic signature; coupled with morpho- logical evidence, this supports a microbial origin of calcite The growth of speleothems such as stalactites and stalag- crystals. SIMS δ13C data suggest that microbially mediated mites through the precipitation of calcite has commonly speleothem formation occurred repeatedly at short intervals been viewed as an abiogenic process (e.g., Kendall and before abiotic precipitation took over. The studied stalactite Broughton, 1978; Broughton, 1983a, b, c). However, there also contains iron and manganese oxides that have been me- is a growing body of research suggesting that microbes diated by microbial activity through extracellular polymeric may play an important role in carbonate precipitation during substance (EPS)-influenced organomineralization processes. speleothem growth (e.g., Jones and Motyka, 1987; Northup The latter reflect paleoenvironmental changes that occurred and Lavoie, 2001; Baskar et al., 2005, 2006; Mulec et al., more than 500 000 yr ago, possibly related to the presence of 2007; Jones, 2010). In caves, a variety of precipitation and a peat bog above the cave at that time. dissolution processes results in the deposition of carbon- Microbial activity can initiate calcite deposition in the ate speleothems, silicates, iron and manganese oxides, sul- aphotic zone of caves before inorganic precipitation of fur compounds, and nitrates, but also in the breakdown of speleothem carbonates. This study highlights the importance limestone host rock. Cave microbes mediate a wide range of of microbially induced fractionation that can result in large destructive and constructive processes that collectively can influence the growth of speleothems and their internal crystal Published by Copernicus Publications on behalf of the European Geosciences Union. 6116 M. Pacton et al.: The role of microorganisms in the formation of a stalactite in Botovskaya Cave, Siberia fabric (Jones, 2010). Constructive processes include microbe calcification, trapping and binding by filamentous microbes, and/or mineral precipitation (Cañaveras et al., 2001; Jones, 2001). Destructive processes include microbially influenced corrosion or dissolution of mineral surfaces that can occur through mechanical attack, secretion of exoenzymes, organic and mineral acids (e.g., sulfuric acid), and a variety of other mechanisms (for a summary please refer to Sand, 1997). Of particular interest in cave dissolution processes are reactions involving iron-, sulfur-, and manganese-oxidizing bacteria (Northup and Lavoie, 2001). Iron oxides and hydroxides are most often observed as coatings or crusts and as powder on clastic cave walls, but they also exist as typical speleothems such as stalactites (e.g., Caldwell and Caldwell, 1980; Jones and Motyka, 1987). Several descriptive studies have estab- lished the association of bacteria with iron deposits in caves, but experimental evidence for an active microbial role in the formation of iron deposits in caves is still lacking (Northup and Lavoie, 2001). Fig. 1. Map of Siberia, showing the location of Botovskaya Cave. With regard to flora and fauna, caves are usually divided into two segments: (i) a twilight zone and (ii) an aphotic zone characterized by no light and few microbes (Jones, 2010). In 2 Study site this study we focus on a stalactite found in the aphotic zone. Botovskaya Cave is located in Siberia (55◦1705900 N, The sample contains mainly calcite and ferromanganese ox- 105◦1904600 E, 750 m above sea level) (Fig. 1 and Fig. S3 in ides, which is unusual in this continental setting as they are Vaks et al., 2013), and developed as a > 68 km-long hori- widely encountered in warm environments (e.g., Spilde et al., zontal maze of passages along tectonic fissures in a 6 to 12 m 2005). The presence of microbes does not automatically im- thick Lower Ordovician limestone layer which is sandwiched ply that they played a role in the formation of the surrounding between marine sandstone and argillite (Filippov, 2000). minerals, because they may simply have been buried dur- The depth of the cave is 40–130 m below the surface. Be- ing mineral precipitation (Polyak and Cokendolpher, 1992; yond a few meters from the small entrances, the light inten- Forti, 2001). Assessment of the exact role that the microbes sity level in the cave is zero. Its passages are characterized played in the mineral precipitation is usually considered im- by massive clay infillings. The cave is located in discontinu- possible to make (Jones, 2010). However, by combining mi- ous permafrost, as evident from massive cave ice bodies with croscopical and geochemical evidence at a high spatial res- < 0 ◦C temperatures in the western part of the cave. Micro- olution, we aim to constrain precisely the role of microbes climatic monitoring reveals that cave air temperature varies in stalactite formation in this cave. Calcite δ13C has been only slightly, from just below 0 ◦C in the cold zones to 1.6– interpreted as reflecting surface vegetation changes (C3 vs. 1.9 ◦C in its warmest parts (see Fig. S4 in Vaks et al., 2013). C4: Brook et al., 1990; Dorale et al., 1992; Bar-Matthews The eastern cave section is the only area where water seepage et al., 1997; Hou et al., 2003; Denniston et al., 2007), al- occurs and speleothems grow today (Vaks et al., 2013). Sam- though more recent studies point to cave air ventilation as a ple SB-p6915 was collected several hundred meters from the major influence (Tremaine et al., 2011). It has been demon- nearest entrance, in the wet and aphotic zone. strated that degassing of CO from the dripwater controls the 2 The region surrounding the cave is covered by taiga rate of calcite precipitation (Mickler et al., 2004; Spötl et al., forest and receives ca. 400 mm annual precipitation. 2005; Bourges et al., 2006; Baldini et al., 2008; Kowalczk Mean annual surface air temperature is −2.8 ◦C, ranging and Froelich, 2010), as well as the isotopic composition of from +35 to −40 ◦C. dripwater and subsequent calcite (Mattey et al., 2008; Müh- linghaus et al. 2007, 2009; Oster et al., 2010; Frisia et al., 2011; Lambert and Aharon, 2011). These studies show that – 3 Methods if carefully evaluated – δ13C can be used as a paleoclimatic indicator. However, the potential impact of microbial activity 3.1 Samples on carbon isotope fractionation is usually overlooked in pa- leoclimate studies. We aim to investigate this impact in detail For this study we use subsamples from stalactite SB-p6915, in the current work. which is asymetrically shaped and has a diameter of 3.5– 4 cm. The sample was found in 2001, more than 1 km deep inside Botovskaya Cave (Supplement Fig. 1), and consists Biogeosciences, 10, 6115–6130, 2013 www.biogeosciences.net/10/6115/2013/ M. Pacton et al.: The role of microorganisms in the formation of a stalactite in Botovskaya Cave, Siberia 6117 3.2 Mineralogy For X-ray diffraction analyses (XRD), samples were ground to a fine powder in an agate mortar. Samples were de- posited on a silicon wafer in a plastic sample holder. We employed a Bruker AXS D8 Advance instrument at ETH Zurich, equipped with a scintillation counter and automatic sampler rotating the sample. 3.3 Microscopy The laminae observed in stalactite SB-p6915 were stud- ied by scanning electron microscopy (SEM) on polished and platinum-coated thin sections, using a Zeiss Supra 50 VP SEM at the University of Zurich, Switzerland. Semi- quantitative elemental analyses of micron-sized spots were obtained using an energy dispersive X-ray spectrometer Fig. 2. Location of the drilling trench of the stalactite from (EDS; EDAX, University of Zurich, Switzerland) during Botovskaya Cave at Layer B (upper arrow), where the sample for U- SEM observations. Th dating was taken. Note the asymmetric distribution of ferroman- ganese oxides in the stalactite. At least 2 hiatuses (marked by ar- rows) divide between Layer B and Hiatus D-E, where the youngest 3.4 Isotope analysis microbial presence can be seen as a thin dark-colored layer.
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